horticulturae
Review
Overview of Melatonin’s Impact on Postharvest Physiology and
Quality of Fruits
Syariful Mubarok 1, * , Erni Suminar 1 , Adzkia Husnul Abidat 2 , Citra Ayu Setyawati 2 , Erik Setiawan 2
and Adine Syabina Buswar 2
Citation: Mubarok, S.; Suminar, E.;
Abidat, A.H.; Setyawati, C.A.;
Setiawan, E.; Buswar, A.S. Overview
of Melatonin’s Impact on Postharvest
Physiology and Quality of Fruits.
Horticulturae 2023, 9, 586.
https://doi.org/10.3390/
horticulturae9050586
Academic Editors: Maribel
Ovando-Martínez, Ramón
Francisco Dórame Miranda
and Carmen Lizette Del Toro
Sánchez
Received: 20 April 2023
Revised: 11 May 2023
Accepted: 14 May 2023
Published: 15 May 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Horticulturae 2023, 9, 586. https://doi.org/10.3390/horticulturae9050586
1 Department of Agronomy, Faculty of Agriculture, Universitas Padjadjaran, Sumedang 45363, JB, Indonesia;
erni.suminar@unpad.ac.id
2 Undergraduate Program of Agrotechnology, Faculty of Agriculture, Universitas Padjadjaran,
Sumedang 45363, JB, Indonesia; adzkia20002@mail.unpad.ac.id (A.H.A.);
citra20002@mail.unpad.ac.id (C.A.S.); erik20001@mail.unpad.ac.id (E.S.);
adine20001@mail.unpad.ac.id (A.S.B.)
* Correspondence: syariful.mubarok@unpad.ac.id
Abstract: Fruits are important horticultural commodities because they provide nutrients that help
human health. Fruits are mostly consumed as fresh products; however, there are challenges in
retaining the freshness, such as the rapid ripening process that triggers fruit deterioration and reduces
fruit quality and nutrient content. The postharvest quality of horticultural crops is affected by pre-and
postharvest treatment. Most farmers use chemical compounds and fungicides to prevent postharvest
damage; however, this results in health hazards and environmental pollution. Melatonin can be
used for maintaining and improving postharvest horticultural crops such as fruits. Melatonin is a
new bioactive compound that is a potent free radical scavenger and antioxidant. It has been studied
as an alternative to harmful chemicals used commercially in the postharvest management of fresh
products. For human health, melatonin plays a regulatory role in circadian and seasonal rhythms,
sleep, retinal functions, and the immune system. In plants, melatonin regulates many biological
processes, particularly when plants have experienced abiotic stress, germination, aging, and growth.
The effect of exogenous melatonin on fruit ripening has focused primarily on the relationship between
melatonin and ethylene plant hormones. Many studies in recent years have discussed melatonin’s
role in plants, particularly in delaying plant aging as an alternative way of increasing fruit shelf
life. This review provides a comprehensive overview of melatonin biosynthesis in plants, factors
that affect the content of melatonin in fruit, melatonin mechanisms in fruit ripening, the impact of
melatonin on postharvest fruit quality, the effect of melatonin on postharvest quality, and the change
in metabolite content of horticultural products, particularly fruits.
Keywords: metabolite; horticulture; antioxidant
1. Introduction
Melatonin is a biomolecule that has pleiotropic effects on living organisms ranging
from the smallest microbes (bacteria) to humans, animals, and plants, but most of it is re-
lated to the regulation of circadian rhythms and biological cycles in animals and plants [1,2].
Melatonin (N-acetyl-5-methoxy tryptamine) was discovered in plants in 1995 and is struc-
turally similar to indole amine compounds such as tryptophan, auxin, and serotonin [1]. In
plants, melatonin functions as a biological signal involved in several plant processes, such
as photosynthesis, development, defense, germination, efflorescence, germination, flow-
ering, leaf senescence, membrane integrity, root development, osmoregulation, and plant
protection against biotic and abiotic stresses [3,4]. For postharvest purposes, melatonin
improves the postharvest quality of horticultural crops [5].
Fruit is a popular horticultural product owing to its good taste and high nutritional
content, such as polyphenolic compounds, organic acids, and vitamins [6]. Fruit is mainly
https://www.mdpi.com/journal/horticulturae
Horticulturae 2023, 9, 586 2 of 15

consumed as a fresh product. However, several problems were found in fruits, such as
perishability and rapid aging during postharvest storage, causing decreased nutritional
content and taste quality, and reduced consumer interest [7]. Moreover, undesirable
changes occurred during fruit ripening, such as softening, shrinkage, ethylene production,
and fruit decay [8–10]. Several methods have been studied to prevent postharvest losses
of perishable products, such as preventing the ethylene effect at the receptor level using
1-methylcyclopropene as an ethylene inhibitor [11–14], mutation techniques in ethylene
receptor genes [15–18], modified atmospheres, and applying chemical compounds, such as
fungicides, waxing, and nitric oxide [19]. However, applying chemical compounds results
in health hazards and environmental pollution; in particular, pesticide residues on fresh
fruits and vegetables will be a potential hazard for consumers. Recently, melatonin has
been studied as an alternative to harmful chemicals used commercially in the postharvest
management of fresh products such as fruits [1].
The postharvest quality of horticultural crops is affected by pre- and postharvest
treatment. Unfavorable conditions during growth and storage will be a major problem
in reducing and losing postharvest quality. Melatonin is a potential nontoxic chemical
compound that can be used for pre- or postharvest treatment. In the field, melatonin
increases pathogen/disease resistance, such as Marssonina apple blotch fungal disease
against Diplocarpon mali [20], inhibits mycelial growth of Phytophthora infestans in potato [21],
increases resistance against Podosphaera xanthii and Phytophthora capsici in cucurbits and
watermelon [22], and increases plant growth and reduces plant stresses, such as salinity,
cold, heat, drought, and heavy metal [1,23]. Preventing unfavorable conditions during
growth and storage will improve the yield, yield quality, and postharvest characteristics of
fresh products.
The effect of melatonin on postharvest quality is still being studied in depth because
the different effects depend on the species. Several studies have reported that melatonin
regulates fruit ripening by regulating ethylene biosynthesis and its roles in harvested prod-
ucts and provide evidence that melatonin is involved in fruit maturation. However, the
effect of melatonin on regulating ethylene biosynthesis that affects fruit ripening differs
depending on the fruit species. In some species, melatonin improves postharvest qual-
ity [24], such as by promoting fruit ripening and improving the fruit quality of tomatoes
and grapes [25,26], or by delaying fruit senescence and increasing the chilling tolerance
of peaches [27]; however, in some species, it reduces postharvest decay and maintains
nutritional quality, such as in strawberries [28]. Previous studies have shown that using
melatonin is important in the postharvest handling of several horticultural crop commodi-
ties and in improving postharvest quality. This review provides a comprehensive overview
of the effect of melatonin on postharvest quality and the change in the metabolite content
of horticultural products, particularly fruits.

2. Melatonin Biosynthesis in Plants

Compared to animals, the biosynthetic process of melatonin in plants is more complex.
Melatonin biosynthesis typically occurs in chloroplasts but can also in mitochondria [1].
Six enzymes are involved in melatonin biosynthesis: L-tryptophan decarboxylase (TDC),
tryptamine 5-hydroxylase (T5H), serotonin N-acetyltransferase (SNAT), acetyl serotonin
O-methyltransferase (ASMT), caffeic acid 3-O-methyltransferase (COMT), and putative
tryptophan hydroxylase (TPH) that has not yet been identified [29].
Figure 1 shows the melatonin biosynthesis. The first step in melatonin biosynthe-
sis process is the conversion of tryptophan, which is the key compound for melatonin
biosynthesis, to serotonin through two different pathways, namely the hydroxylation and
decarboxylation processes. Decarboxylation occurs more commonly in plants compared
to hydroxylation. The first pathway is the decarboxylation of tryptophan to tryptamine
with the help of TDC, then tryptamine is hydroxylated at the 5th position of the indole
ring to form 5-hydroxytryptamine (serotonin) with the help of tryptamine-5-hydroxylase
(T5H); conversely, the second pathway involves the hydroxylation of tryptophan to 5-
3, 9, x FOR PEER REVIEW 3 of 16
Horticulturae 2023, 9, 586 3 of 15

hydroxytryptophan with the help

hydroxytryptophan withof theTPH, followed
help of by the
TPH, followed bydecarboxylation
the decarboxylationprocess of 5-
process of
hydroxytryptophan to form serotonin
5-hydroxytryptophan with thewith
to form serotonin helptheofhelp
TDC [29].[29].
of TDC

Biosynthetic
Figure 1.pathway
Figure 1. Biosynthetic of pathway of melatonin.
melatonin. Solid
Solid lines lines
and and dotted
dotted lineslines indicate
indicate common and
common and al-
alternative pathways of melatonin biosynthesis, respectively. TDC, L-tryptophan
ternative pathways of melatonin biosynthesis, respectively. TDC, L-tryptophan decarboxylase; T5H, decarboxylase;
T5H, tryptamine 5-hydroxylase; TPH, tryptophan hydroxylase; SNAT, serotonin N-acetyltransferase;
tryptamine 5-hydroxylase; TPH, tryptophan hydroxylase; SNAT, serotonin N-acetyltransferase;
ASMT, acetylserotonin O-methyltransferase; COMT, caffeic acid 3-O-methyltransferase.
ASMT, acetylserotonin O-methyltransferase; COMT, caffeic acid 3-O-methyltransferase.
The second step in the melatonin biosynthesis process is the process of converting
The second step into
serotonin melatonin
in the throughbiosynthesis
melatonin two different pathways;
processnamely
is theSNATs catalyze
process of serotonin
converting
into N-acetyl-serotonin, and then, N-acetyl-serotonin is converted into melatonin with the
serotonin into help
melatonin through two different pathways; namely SNATs catalyze seroto-
of ASMTs. The second pathway is the process of converting serotonin into 5-methoxy
nin into N-acetyl-serotonin,
tryptamine with and then,
the help of N-acetyl-serotonin is converted
ASMTs or COMT, followed into melatonin
by the process of convertingwith
the help of ASMTs. The second pathway is the process of converting serotonin into 5-
5-methoxy tryptamine into melatonin catalyzed by SNATs or COMT [29,30].
methoxy tryptamine with the help of ASMTs or COMT, followed by the process of con-
3. Factors That Affect the Melatonin Content in Fruits
verting 5-methoxy tryptamine
3.1. Genetic Makeup into melatonin catalyzed by SNATs or COMT [29,30].
Many studies have broadly examined the presence of melatonin in plants, including
3. Factors Thatvarious
Affectvegetables,
the Melatonin
cereals, Content in and
fruits, seeds, Fruits
medicinal plants [31,32]. The melatonin
content of fruits varies depending on the type and variety of the fruit. Several stud-
3.1. Genetic Makeup
ies have reported endogenous melatonin in different plant families, such as Rosaceae,
Vitaceae,
Many studies have Arecaceae. examined
andbroadly As shown inthe
Table 1, the concentration
presence of endogenous
of melatonin melatonin
in plants, including
varies among species in the same family or in different families, and also according to the
various vegetables, cereals, fruits, seeds, and medicinal plants [31,32]. The melatonin con-
quantification method.
tent of fruits varies depending on the type and variety of the fruit. Several studies have
reported endogenous melatonin in different plant families, such as Rosaceae, Vitaceae, and
Arecaceae. As shown in Table 1, the concentration of endogenous melatonin varies among
species in the same family or in different families, and also according to the quantification
method.

Table 1. Endogenous melatonin in different fruit families.

Horticulturae 2023, 9, 586 4 of 15

Table 1. Endogenous melatonin in different fruit families.

Family Species Concentration (ng/g) References

Actinidiaceae Actinidia chinensis 0.02 [3]
Anacardiaceae Mangifera indica 0.70 [33]
Phoenix dactylifera 0.17 [34]
Arecaceae Phoenix dactylifera 0.17 [34]
Bromeliaceae Ananas comosus (L.) Meri. 0.04 [3]
Ananas comosus (L.) Meri. 0.28 [35]
Ananas comosus (L.) Meri. 0.30 [36]
Ananas comosus (L.) Meri. 0.26 [37]
Ananas comusus 0.04 [38]
Caricaceae Carica papaya 0.24 [33]
Cucurbitaceae Cucumis sativus L. 0.03 [3]
Cucumis sativus L. 0.59 [35]
Cucumis sativa 0.03 [38]
Juglandaceae Juglans regia L. 3.5 [39]
Lythraceae Punica granatum L. 0.17 [35]
Musaceae Musa paradisiaca L. 0.47 [40]
Musa sapientum L. 0.01 [36]
Musa ensete 0.66 [35]
Rosaceae Malus domestica (Borkh) 0.05 [3]
Malus domestica (Borkh) 0.16 [35]
Malus domestica 0.05 [38]
Prunus cerasus L. 18.00 [41]
Prunus avium L. 0.01–0.22 [42]
Prunus avium L. 0.01–124.7 [35]
Prunus cerasus 15.00 [41]
Prunus cerasus 12.3–22.9 [43]
Fragaria ananassa Duch. 0.14 [35]
Fragaria ananassa Duch. 0.01 [3]
Fragaria ananassa Duch. 5.50–11.26 [44]
Fragaria magna 0.01 [38]
Rutaceae Citrus sinensis Osbeck. 0.15 [33]
Solanaceae Solanum lycopersicum 0.5 [40]
Solanum lycopersicum 0.3 [35]
Solanum lycopersicum 7.5–250 [45]
Solanum lycopersicum 0.03 [38]
Solanum lycopersicum 1.2–3.4 [46]
Solanum lycopersicum 4.1–114.5 [44]
Vitaceae Vitis vinifera L. 0.97 [47]
Vitis vinifera L. 1.2–1.50 [48]

3.2. Environment
Another factor that influences the content of melatonin in fruit is the growth envi-
ronment. Based on the research by Burkhardt et al. [41] on Montmorency tart cherries,
no significant difference was noted when planted in different locations. However, this
statement was refuted by the research of Arnao and Hernandez-Ruiz [49], who showed
that environmental conditions affected melatonin levels, which were observed in several
studies of indoor and outdoor tomato cultivation in different environmental conditions.
Melatonin concentration can also be affected by external factors such as stress. Abiotic
stresses can stimulate melatonin synthesis in plants. Melatonin can help ease the negative
consequences, particularly oxidative stress in plants. Melatonin can efficiently cancel the
deleterious effects of toxic oxygen derivatives because of its ability to neutralize them and
increase plant resilience against stress [32]. Moreover, melatonin accumulates in plant tis-
sues as a protective molecule in response to different environmental conditions. Ultraviolet
(UV)-B radiation increased endogenous melatonin up to 7-fold in Glycyrrhyza uralensis com-
pared with the control [50]. The application of zinc salts, hydrogen peroxide, and sodium
chloride in barley and lupin plants resulted in increased endogenous melatonin [36,51].
Horticulturae 2023, 9, 586 5 of 15

Salt stress increased endogenous melatonin up to 2- and 6-fold in the roots and cotyledons
of sunflower seedlings, respectively [52].

3.3. Fruit Ripening Stages of Development

Fruit ripening is another factor that affects the melatonin content of the fruit. According
to research by Murch et al. [53] on berries, the melatonin concentration in all berries
decreases with fruit ripening. Vitalini et al. [54] observed that as the fruit ripens, the
melatonin content in grape skins decreases but increases in the seeds and fruit flesh.
Another study found that cherries with the highest maturity level had the highest melatonin
content [42].

4. Melatonin Mechanism in Fruit Ripening

Ripening is a normal process during fruit maturation. Several biochemical and physi-
ological changes occur, such as those involving flavor, color, aroma, and texture, and there
are also increases in ethylene production and respiration [16,55]. Ethylene is an important
plant hormone for fruit ripening, triggering climacteric fruit ripening [56]. Several studies
have reported that melatonin affects ethylene biosynthesis pathways, but it differs among
species. Melatonin may induce or inhibit ethylene production during different processes in
plants. In tomatoes, melatonin increased ethylene production by upregulating the expres-
sion of 1-aminocyclopropane carboxylic acid synthase 4 (ACS4) [26]. However, in pears,
melatonin reduces ethylene production [57]. In bananas, melatonin can be used to ripen
bananas of different varieties, and it can be used for biological control of postharvest fruit
ripening and quality [58].
The effect of exogenous melatonin on the fruit ripening stage is related to the re-
lationship between melatonin and plant ethylene hormones. Sun et al. [26] found that
exogenous melatonin increased ethylene biosynthesis before and after the climacteric peak
by increasing the expression of 1-aminocyclopropane-1-carboxylic acid (ACC) synthetase
along with the ethylene receptor gene. Proteins that are involved in ripening, cell walls,
carbohydrates, flavonoids, and fatty acid biosynthesis are affected by melatonin. Proteins
are involved in anthocyanin biosynthesis during fruit ripening, showing that exogenous
melatonin regulates fruit ripening and fruit senescence [59]. Strawberries treated with
0.1–1.0 mM melatonin within 5 min and then stored later at 4 ◦ C and 90% humidity showed
a decrease in spoilage and weight [60]. The aging parameters include color, firmness,
total dissolved solid content, titratable acidity, hydrogen peroxide, and malondialdehyde
(MDA). In contrast to the increased phenolic and flavonoid content, a higher antioxidant
capacity was produced. This exogenous melatonin treatment also increased the expression
of endogenous melatonin geosynthetic genes.
Melatonin promotes abscisic acid biosynthesis on various ethylene signaling elements
and increases various primary and secondary metabolic steps to accelerate fruit ripening
by promoting ethylene and anthocyanin biosynthesis. The effect of melatonin on aging
happens in reverse, delaying melatonin through a redox network involving reactive oxygen
species (ROS) and reactive nitrogen species (especially NO), lowering ROS levels, and
upregulating various antioxidant enzymes and metabolites that delay aging. Further-
more, several aging-related genes are derived from melatonin to improve fruit quality and
extend the shelf life. Melatonin treatment has different effects on ethylene biosynthesis.
At one time, it can induce 1-aminocyclopropane-1-carboxylic acid oxidase (ACO) and
1-aminocyclopropane-1-carboxylic (ACC) synthase (ACS) to promote fruit ripening. How-
ever, melatonin also downregulates the same enzymes at other times to reduce ethylene
formation and promote longer shelf life through delayed aging. In this process, ripening
is genetically regulated and characterized by the action of ethylene, whereas aging is an
oxidative process involving damage and disruption. These two processes are different but
inseparable [24,49].
Horticulturae 2023, 9, 586 6 of 15

5. The Effects of Melatonin on Postharvest Damages

Plants need external agents such as melatonin to grow properly from pre- to posthar-
vest. During postharvest, plants are susceptible to pest and disease attacks, which will
result in a decrease in plant quality. Melatonin protects the plants from pests and diseases. It
was reported that the increase in disease resistance might be due to the effects of increasing
endogenous melatonin, inducing salicylic acid (SA) signaling pathways associated with
ROS burst and pathogenesis-related proteins (PRs), activating phenylpropanoid pathways,
and increasing the content of lignin, phenolic compounds, and flavonoids in plants. There-
fore, applying postharvest melatonin is highly recommended because it can be a natural
and economical elicitor in controlling postharvest spoilage; for example, in tomatoes [59]
and strawberries [60] as a valuable approach to increasing fruit quality, and in bananas [58]
and pears [57].
Melatonin treatment increases disease resistance to Pseudomonas syringae pv. on Ara-
bidopsis [61] and against Marssonina blotch (Diplocarpon mali) on apples [33], against
the leaf pathogen Podosphaera xanthii (powdery mildew), and the soil-borne oomycete
Phytophthora capsici in watermelon and other Cucurbita [22]. This result was associated
with increased NO production, which then arranged gene transcription related to SA, acti-
vated cell wall invertase-dependent pathogen defense pathways, maintained intracellular
hydrogen peroxide (H2 O2 ) concentrations, and mediated defense signaling.
Another study also mentioned that melatonin is involved in several plants’ responses
to biotic and abiotic stress. For instance, Lee et al. [62] reported that knockout of serotonin
N-acetyltransferase (SNAT), a rate-limiting enzyme in the melatonin biosynthetic pathway
in Arabidopsis, results in decreased melatonin and SA, which are associated with greater
susceptibility to pathogens. Recently, several studies have indicated that the effect of
melatonin treatment on increasing endogenous melatonin levels is associated with the
increased biosynthetic expression of melatonin genes, thereby contributing to the quality
care of fresh strawberries or the inhibition of browning in longan fruit [60,63].
The presence of melatonin inhibits the development of pathogens such as gray mold
during postharvest. The effect of melatonin on plants makes them resistant to decay and
disease. Endogenous melatonin given to plants receives a burst of ROS, which has relevance
for the upregulation of several genes involved in melatonin biosynthesis. This ROS burst
is induced by chemical elicitors and triggers the activation of a H2 O2 -fueled defense
response, which is highly correlated with lower susceptibility to Trichothecium roseum in
muskmelons [64], Penicillium expansum in apples [65], or Colletotrichum musae infection
in bananas [66]. In addition, SA, as an important endogenous immune signal, triggers
resistance that can be obtained through the systematic induction of a response to the disease
resistance in fruits [67].
Several studies showed that melatonin treatment increased ROS in cherries due to
the formation of O2 , whereas the H2 O2 content increased significantly with melatonin
treatment. Furthermore, melatonin treatment significantly increased endogenous SA con-
tent and the levels of PRs such as CHI and GLU. Moreover, NO is considered an impor-
tant regulatory molecule in the response of plants to pathogen infection in synergy with
ROS [68]. NO interacts with H2 O2 in resistance responses, for example, in tomatoes against
Rhizopus nigricans [69], in peaches against Monilinia fructicola [70], and in oranges against
Colletotrichum gloeosporioides during postharvest [71]. However, not all plants immediately
respond to melatonin postharvest. Likewise, the effect of melatonin application in regulat-
ing fruit development or ripening differs by dose or fruit type or cultivar [72]. Thus, the
effect of melatonin treatment on regulating ROS signaling to activate the SA signaling path-
way may be involved in the disease resistance induced in tomato fruit during postharvest.
Moreover, melatonin has other effects as observed in Fuji apples in cold storage,
which showed that melatonin can reduce ethylene production compared to the control
treatment. Respiration rates immediately decreased in cold storage. The rate of CO2
production gradually decreased from 21 to 35 days of storage, followed by a steady rate
of CO2 production toward the end of storage. The respiration rate pattern of the control
Horticulturae 2023, 9, 586 7 of 15

and melatonin treatments was similar, except that at the end of storage, if melatonin was
not given, ethylene production increased rapidly compared to the melatonin treatment, so
it suppressed ethylene production to a relatively lower level than the fruit control. The
physical impact of receiving melatonin treatment on the study of Fuji apples showed
that after 56 days of fruit storage, the fruit retained its skin appearance as it did during
harvest season compared with the control fruit that had black necrotic spots on the skin.
The underlying metabolic changes between control and melatonin-treated fruit at 56 days
of storage were investigated by UPLC-Q-TOF MS-based metabolic profiles. The results
showed that 40 metabolites, including amino acids, sugars, organic acids, and phenols
were expressed differently in the melatonin-treated samples compared with controls. Most
importantly, the results showed that applying exogenous melatonin increased internal
melatonin levels in apples. Several other metabolites such as phloretin, (+)-catechin,
isoquercitrin, and procyanidins were also influenced by melatonin treatment, which might
positively affect the cold storage of apples [73].

6. Melatonin Effects on Fruits

Many studies have proven that melatonin can effectively influence changes in fruit metabo-
lite content, such as increasing and maintaining ascorbic acid, phenol, and flavonoids [63]. The
application of melatonin changes the physiological index during ripening and postharvest
fruit storage, such as increased phenol, flavonoid, amino acid, fatty acid, and antho-
cyanin [74–76].
Subsequently, melatonin has also been reported as an inducer of nonenzymatic antioxi-
dant accumulation, including phenolics, ascorbic acid, flavonoids, and anthocyanins [77–80].
Melatonin can increase the total phenol and flavonoid content by regulating the cinnamic
acid-4-hydroxylase (C4 H) and phenylalanine ammonia lyase (PAL) activities in the phenyl-
propanoid metabolic pathway, which regulates flavonoid synthesis and other metabo-
lites [78]. Melatonin can increase C4 H activity during initial storage but decrease it during
final storage [78]. This also occurs in PAL activities, where melatonin causes a significant
increase in activity during final storage [78].
In general, during the storage of ascorbic acid, the content decreases due to the
change of L-ascorbic acid to dehydroascorbic acid due to the ascorbic acid oxidase pro-
cess [81]. However, current research indicates that melatonin treatment can change the
ascorbic acid content in fruit, resulting in a higher level of resistance to oxidative stress
during fruit ripening [19,27,82]. Table 2 shows the different responses that may occur
when applying melatonin to fruit due to differences in species, stages of fruit maturity, and
treatment concentrations.
Horticulturae 2023, 9, 586 8 of 15

Table 2. Several studies on the effect of exogenous melatonin on the postharvest treatment of fruits.

Family Species Optimum Concentration (µM) Effects Application References

Delayed the decline of fruit hardness, reduced the loss of soluble
protein, and lowered the accumulation of malondialdehyde; improved
Actinidiaceae Actinidia deliciosa 100 Soaked for 30 min [83]
total phenolic, total flavonoid and flavanol content, and the antioxidant
capacity; and delayed fruit softening.
Maintained firmness, improved ascorbic acid and phenolic compound
content, and antioxidant capacity; controlled the activity of polyphenol
Mangifera indica L. 1000 iImmersed for 10 min [19]
oxidase; and increased the activity of the catalase (CAT) and
peroxidase enzymes during storage.
Reduced respiration rate and ethylene production; increased firmness,
titratable acidity, and ascorbic acid content; lowered weight loss, total
Anacardiaceae soluble solids, pH, total soluble solid and acidity ratio; maintained a
Mangifera indica L. 100 Dipped for 120 min [84]
higher concentration of total phenolics and total flavonoids; increased
the activities of antioxidant enzymes superoxide dismutase and CAT;
and improved membrane stability.
Improved postharvest quality; maintained firmness; improved
Mangifera indica 1000 bioactive compounds; increased phenolic content; and increased the Immersed for 10 min [19]
activity of CAT and peroxidase (POD).
Delayed fruit softening; reduced anthracnose incidence; enhanced
antioxidant activity; and decreased fruit oxidative injury; induced total
phenol, total flavonoid, and ascorbic acid accumulation; increased CAT,
ascorbate peroxidase, NADH oxidase, glutathione reductase,
Caricaceae Carica papaya 400 Immersed for 2 h [85]
polyphenol oxidase, superoxide dismutase, and peroxidase activity;
enhanced the activity of defense-related enzymes, such as chitinase,
4-coumaric acid-CoA-ligase, and phenylalanine ammonia lyase;
repressed lipid metabolism.
Increased chlorophyll, total soluble solids (TSS), soluble sugar, soluble
Cucurbitaceae Momordica charantia L. 120 protein, and ascorbic acid (AsA); promoted the synthesis of total Immersed for 10 min [86]
phenols and flavonoids.
Maintained the contents of AsA, anthocyanin, and total
phenol; reduced the accumulation of reactive oxygen species and
Ericaceae Vaccinium spp. 50 membrane lipid peroxidation; and promoted the activities of ascorbate Sprayed [87]
peroxidase (APX), glutathione S-transferase, and phenylalanine
ammonia lyase (PAL).
Horticulturae 2023, 9, 586 9 of 15

Table 2. Cont.

Family Species Optimum Concentration (µM) Effects Application References

Musaceae Musa paradisiaca L. 200–500 Inhibited the ripening process. Soaked for 2 h [88]
Maintained postharvest quality, delayed fruit softening, and reduced
the incidence of anthracnose; enhanced the antioxidant capacity and
Myrtaceae Psidium guajava L. 600 reduced the oxidative damage; enhanced the activities of CAT, Soaked for 2 h [78]
superoxide dismutase, ascorbate peroxidase, and glutathione
reductase; and maintained total flavonoids and AsA.
Reduced ethylene production and weight loss; maintained skin
Malus domestica Borkh 1000 structure; increased the activity of three enzymes, including POD, Sprayed [89]
superoxide dismutase (SOD), and CAT.
Malus domestica Borkh 200 Inhibited gray mold diseases. Immersed [90]
Reduced weight loss, decay incidence, and respiration rate; maintained
firmness, TSS, and AsA contents; enhanced the activities of superoxide
Prunus persica 100 dismutase, CAT, peroxidase, and ascorbate peroxidase; decreased the Dipped for 10 min [27]
activity of lipoxygenase, levels of superoxide anion and hydrogen
peroxide, and malondialdehyde content.
Reduced chilling injury; increased extractable juice rate and TSS;
Prunus persica 100 enhanced expression of PpADC, PpODC, and PpGAD; increased Immersed for 2 h [73]
polyamines and γ-aminobutyric acid (GABA) contents.
Melatonin-treated nectarines exhibited higher total antioxidant activity
than controls, which was correlated primarily to the increase in the
Rosaceae levels of total phenolics and to a decrease in the loss to AsA and
Prunus persica 1000 flavonoids contents. These results demonstrated that melatonin Immersed for 30 min [91]
treatment could be a good practice for extending postharvest
life of nectarine fruits, maintaining the appearance and nutrient value,
and reducing the loss of health-promoting compounds.
Reduced weight loss and maintaining greater firmness; increased AsA
Prunus salicina 1000 Immersed [77]
content, total phenolic content, and antioxidant; reduced decay rate.
Prunus salicina 100 Enhanced S-adenosylmethionine decarboxylase and TGase activities. Immersed for 100 min [92]
Reduced decay and weight loss of fruit; delayed senescence; reduced
the accumulation of hydrogen peroxide (H2 O2 ) and malondialdehyde
(MDA); increased the total phenolics and flavonoid contents resulting
Fragaria × ananassa,
100 or 1000 in the higher antioxidant capacity, extending the postharvest life, and Immersed for 5 min [60]
Duch.
improving the fruit quality; enhanced the expression of melatonin
biosynthetic genes including FaTDC, FaT5H, FaSNAT, and FaASMT;
and increased the content of endogenous melatonin.
Fragaria × ananassa, Increased the acumulation of H2 O2 ; increased the activity of SOD;
100 Immersed for 2 h [93]
Duch reduced the activity of CAT and APX; and decreased the fruit decay.
Horticulturae 2023, 9, 586 10 of 15

Table 2. Cont.

Family Species Optimum Concentration (µM) Effects Application References

Reduced flesh browning and decay incidence; increased phenols,
Prunus avium L. 100 flavonoids, anthocyanins accumulation, and antioxidant potential; Dipped for 5 min [94]
maintained the membrane integrity.
Inhibited weight loss and firmness; increased sugar, acid, and
Eriobotrya japonica Lindl. 50 phytochemical content; decreased MDA content; improved antioxidant Immersed for 30 min [95]
capacity; and reduced lignin content.
Delayed the ethylene production; increased fruit firmness; increased
Pyrus communis L. 100 Immersed for 12 h [57]
antioxidant; and delayed the senescene process.
Decreased respiration and weight loss rates; increase fruit firmness,
TSS, soluble sugar content, titratable acidity, and citrus color index;
impeded the accumulation of hydrogen peroxide and
Citrus sinensis (L.)
Rutaceae 200 malondialdehyde; inhibited reactive oxygen species (ROS) burst and Dipped for 5 min [96]
Osbeck
oxidative damage; increased the activity and expressions of CAT,
superoxide dismutase, ascorbate peroxidase, and glutathione
reductase; and delayed postharvest senescence.
Enhanced degradation of pesticides, such as chlorothalonil, malathion,
Rhamnaceae Ziziphus jujuba Mill. 100 and glyphosate; delayed fruit senescence by reduced weight loss, Sprayed [97]
decay incidence, and firmness.
Delayed the development of pericarp browning, inhibited lipid
Litchi chinensis Sonn. 400 Immersed for 5 min [75]
Sapindaceae degradation, and maintained membrane integrity and energy status.
Inhibited pericarp browning; reduced discoloration during storage;
Litchi chinensis Sonn. 400 Immersed for 5 min [98]
decreased membrane relative leakage rate.
Increased the content of total flavonoids, total phenolics, and lignin;
inhibited gray mold development, induced a ROS burst, increased
endogenous melatonin and salicylic acid (SA); enhanced activities of
Solanum lycopersicum 100 Dipped 60 min [99]
chitinase (CHI) and β-1,3-glucanase (GLU); increased the activities of
PAL, 4-coumarate-coenzyme A ligase (4CL), and POD; and increased
Solanaceae total phenols, flavonoids, and lignin in tomato.
Increased lycopene and water loss; enhanced fruit softening; increased
water-soluble pectin and decreased protopectin; increased ethylene
Solanum lycopersicum 50 Immersed for 2 h [26]
production, accelerated the climacteric phase, enhanced fruit ripening,
and improved quality of tomatoes.
Horticulturae 2023, 9, 586 11 of 15

In papayas, applying melatonin can increase flavonoid and phenolic total content and
ascorbic acid during fruit ripening to increase the accumulation of metabolite content [85].
In guava, melatonin treatment resulted in increased ascorbic acid content accumulation.
Melatonin has the effect of increasing the respiration rate of fruit during final storage, so
this impacts the secondary metabolite content accumulation of guava.
Melatonin application on strawberries can reduce malondialdehyde (MDA) and H2 O2
concentrations while increasing phenolic and flavonoid total content, thereby contributing
to the fruit’s potential for high antioxidant content [60]. However, melatonin treatment
on strawberries resulted in lower ascorbic acid content. The response differed from that
of melatonin treatment in peaches, which showed that it maintained its ascorbic acid
content [27,60]. This may be due to differences in species, fruit conditions, and melatonin
concentrations.
Melatonin treatment of cold-stored fruits with adequate supplies of intercellular
NADPH can result in higher metabolite content. In addition to melatonin treatment in
pomegranates, these conditions activate the ROS prevention system and increase phenyl-
propanoid metabolic pathway activity, resulting in a higher accumulation of phenols and
anthocyanins [28]. PAL activity increased in cherry tomatoes due to melatonin application;
this response positively impacted fruit phenolic and flavonoid total and lignin content dur-
ing storage [99]. In addition, changes in metabolite content have also increased in several
other fruits that have been studied, such as plums [77], mangoes [19,84], and kiwis [83].

7. Conclusions
The primary function of melatonin in fruit is its ability to induce parthenocarpy. It also
promotes ripening and aging of fruits by upregulating and downregulating many gene
elements related to ethylene, anthocyanins, flavonoids, cell wall enzymes, senescence, car-
bohydrate metabolism. Melatonin has the potential to serve as a natural postharvest alter-
native for enhancing postharvest quality, reducing the incidence of spoilage, and increasing
the antioxidant potential of fruits by either increasing or maintaining bioactive compounds.

Author Contributions: Conceptualization, S.M. and E.S. (Erni Suminar); methodology, S.M., A.H.A.,
C.A.S., E.S. (Erik Setiawan) and A.S.B.; validation, S.M.; investigation, A.H.A., C.A.S., E.S. (Erik Seti-
awan) and A.S.B.; resources, S.M., A.H.A., C.A.S., E.S. (Erik Setiawan) and A.S.B.; writing—original
draft preparation, S.M., A.H.A., C.A.S., E.S. (Erik Setiawan) and A.S.B.; writing—review and editing,
S.M. and E.S. (Erni Suminar); visualization, S.M.; supervision, S.M.; project administration, S.M.; fund-
ing acquisition, S.M. All authors have read and agreed to the published version of the manuscript.
Funding: This research received no external funding.
Data Availability Statement: The data are included in the article.
Acknowledgments: We thank all of the members of our laboratory for helpful discussions throughout
the work. We also thank Universitas Padjadjaran, Indonesia, for supporting the APC.
Conflicts of Interest: The authors declare no conflict of interest.

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